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Although studying<jats:italic>F. tularensis<\/jats:italic>pathogenesis is essential to comprehending disease, mammalian infection is just one step in the ecology of<jats:italic>Francisella<\/jats:italic>species.<jats:italic>F. tularensis<\/jats:italic>has been isolated from aquatic environments and arthropod vectors, environments in which chitin could serve as a potential carbon source and as a surface for attachment and growth. We show that<jats:italic>F. tularensis<\/jats:italic>subsp.<jats:italic>novicida<\/jats:italic>forms biofilms during the colonization of chitin surfaces. The ability of<jats:italic>F. tularensis<\/jats:italic>to persist using chitin as a sole carbon source is dependent on chitinases, since mutants lacking<jats:italic>chiA<\/jats:italic>or<jats:italic>chiB<\/jats:italic>are attenuated for chitin colonization and biofilm formation in the absence of exogenous sugar. A genetic screen for biofilm mutants identified the Sec translocon export pathway and 14 secreted proteins. We show that these genes are important for initial attachment during biofilm formation. We generated defined deletion mutants by targeting two chaperone genes (<jats:italic>secB1<\/jats:italic>and<jats:italic>secB2<\/jats:italic>) involved in Sec-dependent secretion and four genes that encode putative secreted proteins. All of the mutants were deficient in attachment to polystyrene and chitin surfaces and for biofilm formation compared to wild-type<jats:italic>F. novicida<\/jats:italic>. In contrast, mutations in the Sec translocon and secreted factors did not affect virulence. Our data suggest that biofilm formation by<jats:italic>F. tularensis<\/jats:italic>promotes persistence on chitin surfaces. Further study of the interaction of<jats:italic>F. tularensis<\/jats:italic>with the chitin microenvironment may provide insight into the environmental survival and transmission mechanisms of this pathogen.<\/jats:p>","DOI":"10.1128\/aem.02037-09","type":"journal-article","created":{"date-parts":[[2009,12,1]],"date-time":"2009-12-01T02:59:47Z","timestamp":1259636387000},"page":"596-608","update-policy":"https:\/\/doi.org\/10.1128\/asmj-crossmark-policy-page","source":"Crossref","is-referenced-by-count":64,"title":["Contributions of<i>Francisella tularensis<\/i>subsp.<i>novicida<\/i>Chitinases and Sec Secretion System to Biofilm Formation on Chitin"],"prefix":"10.1128","volume":"76","author":[{"given":"Jeffrey J.","family":"Margolis","sequence":"first","affiliation":[{"name":"Department of Microbiology and Immunology, Stanford University School of Medicine, Stanford, California 94305"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sahar","family":"El-Etr","sequence":"additional","affiliation":[{"name":"Bioscience and Biotechnology Division, Lawrence Livermore National Laboratory, Livermore, California 94550"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lydia-Marie","family":"Joubert","sequence":"additional","affiliation":[{"name":"Cell Sciences Imaging Facility, Stanford University School of Medicine, Stanford, California 94305"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Emily","family":"Moore","sequence":"additional","affiliation":[{"name":"Department of Microbiology and Molecular Biology, Brigham Young University, Provo, Utah 84602"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Richard","family":"Robison","sequence":"additional","affiliation":[{"name":"Department of Microbiology and Molecular Biology, Brigham Young University, Provo, Utah 84602"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Amy","family":"Rasley","sequence":"additional","affiliation":[{"name":"Bioscience and Biotechnology Division, Lawrence Livermore National Laboratory, Livermore, California 94550"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alfred M.","family":"Spormann","sequence":"additional","affiliation":[{"name":"Department of Civil and Environmental Engineering, Stanford University, Stanford, California 94304"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Denise M.","family":"Monack","sequence":"additional","affiliation":[{"name":"Department of Microbiology and Immunology, Stanford University School of Medicine, Stanford, California 94305"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"235","reference":[{"key":"e_1_3_2_2_2","doi-asserted-by":"publisher","DOI":"10.1128\/AEM.65.9.4285-4287.1999"},{"key":"e_1_3_2_3_2","doi-asserted-by":"publisher","DOI":"10.3201\/eid0707.017740"},{"key":"e_1_3_2_4_2","doi-asserted-by":"publisher","DOI":"10.1111\/j.1462-2920.2007.01491.x"},{"key":"e_1_3_2_5_2","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.mi.22.100168.000511"},{"key":"e_1_3_2_6_2","doi-asserted-by":"publisher","DOI":"10.1016\/S0021-9258(18)54225-3"},{"key":"e_1_3_2_7_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.jmb.2004.05.028"},{"key":"e_1_3_2_8_2","doi-asserted-by":"publisher","DOI":"10.1046\/j.1365-2958.2002.03091.x"},{"key":"e_1_3_2_9_2","doi-asserted-by":"publisher","DOI":"10.1128\/IAI.00430-08"},{"key":"e_1_3_2_10_2","doi-asserted-by":"publisher","DOI":"10.1016\/0092-8674(90)90111-Q"},{"key":"e_1_3_2_11_2","doi-asserted-by":"publisher","DOI":"10.1007\/s12098-008-0180-9"},{"key":"e_1_3_2_12_2","doi-asserted-by":"publisher","DOI":"10.1016\/S0076-6879(99)10004-1"},{"key":"e_1_3_2_13_2","doi-asserted-by":"publisher","DOI":"10.1126\/science.284.5418.1318"},{"key":"e_1_3_2_14_2","unstructured":"Cowan and Steel's manual for the identification of medical bacteria 2004"},{"key":"e_1_3_2_15_2","doi-asserted-by":"publisher","DOI":"10.1038\/417243a"},{"key":"e_1_3_2_16_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.micinf.2003.09.002"},{"key":"e_1_3_2_17_2","doi-asserted-by":"publisher","DOI":"10.1016\/S0014-2565(01)70854-5"},{"key":"e_1_3_2_18_2","doi-asserted-by":"crossref","first-page":"415","DOI":"10.4049\/jimmunol.87.4.415","volume":"87","year":"1961","unstructured":"Eigelsbach, H. 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